Animal Behaviour 80 (2010) 3e8

Contents lists available at ScienceDirect

Animal Behaviour
journal homepage: www.elsevier.com/locate/anbehav

Essay

The central importance of information in studies of animal communication

Robert M. Seyfarth a, *, Dorothy L. Cheney b, Thore Bergman c, d, Julia Fischer e, Klaus Zuberbühler f,
Kurt Hammerschmidt e
a
Department of Psychology, University of Pennsylvania, Philadelphia, PA, U.S.A.
b
Department of Biology, University of Pennsylvania, Philadelphia, PA, U.S.A.
c
Department of Psychology, University of Michigan, Ann Arbor, MI, U.S.A.
d
Department of Ecology & Evolutionary Biology, University of Michigan, Ann Arbor, MI, U.S.A.
e
Cognitive Ethology Laboratory, German Primate Center, University of Gottingen, Gottingen, Germany
f
Wissenschaftskolleg, Institute for Advanced Study, Berlin, Germany

a r t i c l e i n f o
The concept of information plays a central role in studies of animal communication. Animals’ responses
Article history: to the calls of different individuals, to food calls, alarm calls, and to signals that predict behaviour, all
Received 17 December 2009 suggest that recipients acquire information from signals and that this information affects their response.
Initial acceptance 4 March 2010 Some scientists, however, want to replace the concept of information with one based on the ‘manipu-
Final acceptance 26 March 2010 lation’ of recipients by signallers through the induction of nervous-system responses. Here we review
Available online 21 May 2010 both theory and data that argue against hypotheses based exclusively on manipulation or on a fixed,
MS. number: AE-09-00802 obligatory link between a signal’s physical features and the responses it elicits. Results from dozens of
studies indicate that calls with ‘arousing’ or ‘aversive’ features may also contain information that affects
Keywords: receivers’ responses; that acoustically similar calls can elicit different responses; acoustically different
animal communication
calls can elicit similar responses; and ‘eavesdropping’ animals respond to the relationship instantiated by
animal signal
signal sequences. Animal signals encode a surprisingly rich amount of information. The content of this
information
information transmission
information can be studied scientifically.
language ! 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
meaning

The concept of information has played a long and productive role
in the study of animal communication. Empirical research has
attempted to specify the kinds and amount of information trans-
ferred in signalling systems as disparate as the dance language of
honeybees (von Frisch 1967; Seeley 1997), the claw-waving displays
of crustaceans (Dingle 1969), the songs of birds (Vehrencamp 2000),
and the alarm calls of primates (Cheney & Seyfarth 1990). Theo-
retical analyses have relied heavily on the concept of information in
contexts ranging from aggression to courtship to cooperation
(Enquist 1985; Grafen 1990; Maynard Smith 1991).
Nevertheless, this approach has been criticized by a minority of
researchers. Dawkins & Krebs (1978, page 309), for example, argued
that animal signals should be viewed not in terms of information
but in terms of the manipulation of receiver behaviour, arguing that
‘it is probably better to abandon the concept of information transfer
altogether’. The call to abandon information in favour of ‘manipu-
lation’ or ‘assessment/management’ was later renewed by Owings
& Morton (1997, 1998), who suggested that ‘the informational
* Correspondence: R. M. Seyfarth, Department of Psychology, University of
Pennsylvania, 3720 Walnut Street, Philadelphia, PA, U.S.A.
E-mail address: seyfarth@psych.upenn.edu (R.M. Seyfarth).
perspective is not adequate as a concept or methodology to
understand either the evolution or the process of vocal commu-
nication’ (1998, page ix). Along with others, they advocate a view of
communication as management by signallers of the behaviour of
receivers (see also Owren & Rendall 1997, 2001; Owren 2000;
Rendall et al. 2009). For example, following Morton (1977),
Rendall et al. (2009) note that the squeaks, shrieks and screams
of many animals have ‘sharp onsets, dramatic frequency and
amplitude fluctuations, and chaotic spectral structures, which are
exactly the sorts of features that have direct impact on animals’
nervous systems’ (page 236). Similar generalizations hold for alarm
calls which have evolved, they believe, to ‘induce nervous-system
responses’ in receivers (Owren & Rendall 2001, page 61). Finally, the
critics also argue that using terms like information implicitly
commits scientists to the use of human communication, particu-
larly language, as a model for communication in animals. It is
therefore both anthropomorphic and inaccurate (Owings & Morton
1997, 1998).
These opposing views find parallels in studies of animal learning.
For years, behaviourists argued that the mental activities of animals
were not appropriate topics for research, either because they could
not be studied scientifically (methodological behaviourism) or
because they did not exist (radical behaviourism; Skinner 1974).

0003-3472/$38.00 ! 2010 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.anbehav.2010.04.012
4 R.M. Seyfarth et al. / Animal Behaviour 80 (2010) 3e8
Today, however, this view has largely been discarded. Modern
theories of learning have a strong cognitive component, with many
experiments designed to examine the content of animals’
knowledge and the information that animals acquire as a result of
experience (e.g. Colwell & Rescorla 1985; Kamil 1987; Rescorla
1988; Lieberman 2003).
CLARIFYING THE TERMINOLOGY
Current accounts of information are built on the theoretical
advances made by Shannon (1948) and Wiener (1961). Shannon
viewed information as a statistical measure of uncertainty, allowing
for mathematical analyses of information processing. While infor-
mation theory was initially developed to describe information
transmission in technical systems, it quickly found its way into
a range of other disciplines, including psychology and animal
behaviour (e.g. Dingle 1969; Beecher 1989). For our purposes,
treating information as a reduction of uncertainty in the recipient is
useful because it connects communication to learning theory and to
research on the mechanisms by which animals associate signals (or
cues) with each other or with the outcomes of specific behaviours.
INFORMATION IN ANIMAL COMMUNICATION
Whenever there is a predictable relation between a particular
signal and a specific social situation, the signal can be used by
listeners to predict current states or upcoming events; that is, to
provide information. A light that predicts shock, an alarm call that
predicts the presence of an eagle, or a scream that predicts that
a specific individual is involved in a fight all have the potential to
provide a listener with information if they are reliably associated
(Rescorla 1988) with a narrow range of events (Seyfarth & Cheney
2003). In each case we assume that the listener has acquired the
contingent relation between two stimuli and thus reduced its
uncertainty (or gained information) about events in the world: the
light predicts shock, not food; the alarm call predicts an eagle, not
a leopard; the screams predict that one individual, but not another,
is involved in a dispute. We also assume that learning such
associations (if learning is required) is adaptive because it allows
the receiver to predict what is likely to happen next.
Empirical support for this use of information is widespread in
studies of animal communication. It is now clear that individuals in
many species consistently use specific signals in particular social or
ecological contexts and that receivers have learned or otherwise
acquired these contingent relations, gaining information as a result.
For example, honeybees, Apis mellifera, acquire information about
the location of food by observing the details of a worker bee’s
dance (e.g. Seeley 1997). In hermit crabs, Pagurus bernhardus
(Laidre 2009), swamp sparrows, Melospiza georgiana (Ballentine
et al. 2008), and banded wrens, Pheugopedius pleurostictus
(Vehrencamp et al. 2007), certain visual or vocal displays are reli-
able predictors of an individual’s subsequent aggressive behaviour,
and recipients respond as if they know this relation. The begging
calls of cliff swallows, Hirundo pyrrhonota, are individually
distinctive and provide parents with information about individual
identity; the begging calls of barn swallows, H. rustica, do not
(Medvin et al. 1993). In black-capped chickadees, Poecile atrica-
pillus, acoustic features of the ‘seet’ and ‘chick-a-dee’ alarm calls are
correlated with both the type of predator present and the degree of
danger. Playback experiments indicate that listeners acquire this
information from the calls (Templeton et al. 2005). A similar
generalization holds for the alarm calls of African suricates, Suricata
suricatta. Suricates give acoustically different alarm calls to
different predators (jackals, eagles and snakes), and within each
call type produce calls with graded acoustic variation that is
correlated with the urgency of the danger. The suricates’ responses
to call playbacks suggest that, upon hearing an alarm call,
individuals acquire information about both predator type and
urgency (Manser et al. 2001a, b). In California ground squirrels,
Spermophilus beecheyi, acoustically different alarm calls encode
information about urgency, but not predator type (Owings &
Hennessy 1984); in primates, they encode information about
different predators (vervet monkeys, Chlorocebus aethiops: Seyfarth
et al. 1980; Diana monkeys, Cercopithecus diana: Zuberbühler et al.
1999). Macedonia & Evans (1993) discuss the evolution of alarm call
systems that encode different types of information. Finally, in
addition to their acoustically different alarm calls for ground and
aerial predators (Evans et al. 1993), domestic chickens, Gallus gallus
domesticus, produce food calls that signal the presence of food.
Once again, playback experiments indicate that listeners acquire
this information from the calls (Evans & Evans 1999, 2007).
Contrary to the critics’ argument, investigators in these studies
have not naïvely applied linguistic concepts to their subjects, nor
have they claimed that their results demonstrate the same kind of
information transfer found in language. Instead, they have simply
tried to determine whether particular signals predict something
about the world (e.g. the presence of food, the caller’s identity,
a particular kind of predator, or the urgency of danger) or about the
signaller’s next behaviour. In this respect, the critics have set up
a straw man: although information plays a central role in studies of
animal communication, ethology is in no danger of being taken
over by linguistics.
The ‘manipulative’ approach also leaves many interesting ques-
tions unanswered. For example, some of the alarm calls mentioned
above have acoustic features that are arousing and attention getting.
Why, then, are the calls within each species acoustically so
different? Why do individuals respond to them in such different
ways? And why is there such wide variation across species in the
acoustic properties of alarm calls? Rats’ alarms have a whistle-like
structure (Litvin et al. 2007), those of shifaks are low-frequency roar
grunts (Fichtel & Kappeler 2002), while antelope produce snorts
(Tilson & Norton 1981). An exclusive focus on the calls’ arousing
physical characteristics cannot answer these questions.
Nor does the presence of particular acoustic features preclude
the acquisition of information by listeners. Primate screams, for
example, may have ‘aversive’ acoustic qualities, but this does not
preclude recipients from acquiring information from them. In fact,
research on many primates has shown that screams are individu-
ally distinctive (e.g. Hammerschmidt & Fischer 1998). Their
acoustic features can also be correlated with different types of
aggression (Gouzoules et al. 1984), the caller’s role in the interac-
tion (Slocombe & Zuberbühler 2005), or the presence of a particular
‘audience’ (Slocombe & Zuberbühler 2007). As a result, screams
used in playback experiments elicit different responses from
different individuals, or from the same individual under different
circumstances (Cheney & Seyfarth 1980; Gouzoules et al. 1984;
Palombit et al. 1997; Fischer 2004; Fugate et al. 2008; Slocombe
et al. 2009). Here again, an exclusive focus on the screams’
aversive qualities cannot explain these results.
Theoretical Limitations
The critics argue that signals have evolved to manipulate
listeners’ behaviour, and that the acoustic properties of signals take
the form that they do because they have a ‘direct effect’ on listeners’
nervous systems, effects that are difficult for listeners to resist. This
explanation assumes that listeners are automata that can be
manipulated to respond in ways beneficial to the signaller as long
as the right nervous-system buttons are pushed. However, as
Searcy & Nowicki (2005, page 8) point out, ‘The critical flaw with
 R.M. Seyfarth et al. / Animal Behaviour 80 (2010) 3e8
this reasoning is that it does not explain why the receiver would be
selected to respond to the signal at all’. An analysis that allows the
signaller’s behaviour to evolve but does not permit any evolution in
receiver response does not make sense. ‘If there is, on average, no
information of benefit to the receiver in a signal, then receivers
should evolve to ignore that signal. If receivers ignore the signal,
then signalling no longer has any benefit to the signaller, and the
whole communication system should disappear’ (Searcy & Nowicki
2005, page 8). In fact, receivers should evolve responses to signals
only when it is advantageous to do so. And if it does not benefit
receivers to respond in a particular way to a specific acoustic
feature then selection will favour receivers that attend to some
other cue.
Imagine, for example, a species of frog in which males compete
with vocal displays and calls with a low dominant frequency induce
a ‘nervous-system response’ that causes recipients to retreat. This
response will be maintained by natural selection only if it benefits
the recipient. This could happen if calls with a low dominant
frequency accurately predict the caller’s competitive ability,
thereby providing recipients with information about their likely
success if they compete with that opponent. If calls with a low
dominant frequency do not allow accurate assessment, then
selection will favour recipients who base their decision on some
other cue, regardless of any impact that low dominant frequency
might have on their nervous system.
Receivers are not, then, prisoners of the influence that specific
acoustic properties have on their sensory systems. Instead, selection
will favour receivers that act selfishly, adjusting their ‘assessment
rule’ (Grafen 1990, page 521) so that it is most effective in reducing
uncertainty, or in providing them with information.
Finally, Grafen (1990), Searcy & Nowicki (2005), and others have
used the logic of information to explain the evolution of all animal
signals, regardless of modality. This generality constitutes a strength
of their approach. If the explanation based on manipulation is to be
equally general, its advocates will need to show that, just as some
auditory signals like screams manipulate listeners through ‘loud
bursts of jarring broadband noise’ (Rendall et al. 2009), certain
visual signals are equally aversive and manipulative by virtue of
their physical properties. This seems unlikely.
Receiver Flexibility
Supporting the view that selection can act just as forcefully on
receivers as it does on signallers, there is ample evidence that
receivers can learn to respond in specific ways to signals regardless
of the signals’ acoustic properties (think, for example, of learning
experiments using pure tones, white noise, or more naturalistic
vocalizations as stimuli), and that receiver responses can evolve
over time (cases of character displacement, for example, suggest
that receiver response may be more malleable than signal form).
Other studies suggest that there is no obligatory relation
between acoustic features and response. In songbirds that sing more
than one song type, the rate of song type switching increases with
increasing aggression in some species but decreases with increasing
aggression in others (Vehrencamp 2000). Song type switching has
no inherent, immutable effect on listeners’ behaviour. Instead,
listeners attend to song switching rate because it predicts the
singer’s behaviour and thus provides useful information.
Of course, there are also many cases in which acoustic features
are closely linked to call function. Two examples are the relationship
between call frequency and body size in frogs and toads (reviewed
in Searcy & Nowicki 2005), and between formant spacing and body
size in several mammals (reviewed in Fitch & Hauser 2003). In both
of these cases, however, listeners attend to a crucial acoustic feature
not just because it induces a ‘nervous-system response’ (although it
5
may do this) but also because it reduces uncertainty (that is,
provides information) about a competitor.
Finally, Rendall et al. (2009, page 236) argue that ‘a frustrated
primate weanling cannot force its mother to nurse, but can readily
elicit such behaviour with sounds’ that ‘with repetition become
very aversive’. But studies of primate mothereinfant interactions
show that aversive sounds do not always have this effect. Infants do
get weaned, in part because mothers cease responding to their calls
and reject their attempts to nurse (e.g. Hinde 1978; Altmann 1980,
page 175e177; Maestripieri 2002; see also Hammerschmidt et al.
1994).
Calls with Similar Acoustic Features Can Elicit Different Responses
Animals often respond differently to acoustically similar calls.
Chickens, for example, respond in very different ways to food calls
and to ground predator alarm calls even though the calls have
similar acoustic characteristics (Evans & Evans 1999). They also
respond differently to the same food call depending on whether
they already know about the presence of food (Evans & Evans
2007). Upon hearing a vervet monkey’s eagle alarm call, nearby
animals on the ground look up or run into a bush. Animals in a tree
look up and/or run down out of the tree and into a bush, and
animals already in bushes typically do nothing (Seyfarth et al. 1980;
see also Fischer et al. 2000). Listeners’ responses to baboon grunts
depend on both the type of grunt given and details of the social and
ecological context (Rendall et al. 1999; Rendall 2003), including the
recipient’s prior interaction with the caller (Cheney & Seyfarth
1997), and the relation between the caller and individuals with
whom the recipient has recently interacted (Wittig et al. 2007).
These differences in response to the same or acoustically similar
calls cannot be attributed to acoustic features alone, but they are
consistent with the hypothesis that responses depend upon the
integration of information acquired from calls and other contextual
cues.
Calls with Different Acoustic Features Are Judged To Be Similar
Diana monkeys that typically respond to the growl of a leopard
by giving their own leopard alarm call will not give such calls in
response to the growl if they have already heard a Diana’s leopard
alarm call coming from the same area. However, they respond in
typical fashion to the leopard’s growl if they previously heard
a Diana’s eagle alarm call coming from the same area. Although
a leopard’s growl and Diana monkey leopard alarm calls are very
different acoustically, the monkeys respond as if they judge them to
be similar (similar, at least, in the sense that responding to one call
type produces habituation to the other; Zuberbühler et al. 1999). An
exclusive focus on call acoustics cannot explain these results. They
are, however, consistent with the hypothesis that listeners acquire
information from a call, store it in memory, and respond to
subsequent vocalizations depending on some combination of
acoustic features, information provided by the current vocalization
and context, and information stored in memory (for similar results
see Fischer 1988; Cheney & Seyfarth 1988; Rendall et al. 1996;
Hauser 1998).
Animals’ Responses Depend on the Relationship Instantiated by Two
Signals
Song sparrows, Melospiza melodia, sometimes match a neigh-
bour’s song by selecting from among their own repertoire the song
type that most closely matches the song their neighbour just sang
(Beecher et al. 1996). In studies using interactive playback experi-
ments, subjects responded more strongly to a match than
6 R.M. Seyfarth et al. / Animal Behaviour 80 (2010) 3e8
a nonmatch (Burt et al. 2001). It was not just the acoustic properties
of the song that determined the neighbour’s behaviour (although
these were clearly important), but whether or not the two song
types matched (see also Beecher & Campbell 2005).
Territorial songbirds learn to recognise their neighbours and
associate them with specific areas by listening to their songs. They
respond more strongly to a given song type when it comes from an
unfamiliar area than when it comes from a familiar area (Brooks &
Falls 1975; see also Herbinger et al. 2009). When a listener responds
differently to the same vocalization played from different locations,
it integrates information contained in the call itself with informa-
tion stored in memory about the caller’s typical location. The direct
effects of acoustic features alone cannot explain this behaviour.
Songbirds also learn about the competitive abilities of potential
intruders by listening to their singing bouts with the territory
holders’ neighbours (Peake et al. 2002). Such ‘eavesdropping’ is
widespread among animals (McGregor 2005). When a bird or
primate responds differently depending upon whether it has heard
A dominate B or B dominate A (Peake et al. 2002; Bergman et al.
2003), its response depends not just on the calls’ acoustic proper-
ties but also on the relationship between callers. As McGregor &
Dabelsteen (1996, page 416) put it, the eavesdropper ‘gains infor-
mation from an interaction that could not be gained from a signal
alone’. Once again, an exclusive focus on acoustic properties is
insufficient to explain these results.
USING ‘INFORMATION’ IN ANIMAL COMMUNICATION
The experiments cited above, and many others, suggest that the
manipulative hypothesis is insufficient to explain all that happens
when one individual signals to another. Instead, results suggest that
when recipients perceive a signal they acquire information, and the
acquisition of this information (among other things) changes their
behaviour. The information that receivers acquire has content, and
this content can be studied scientifically.
Used in this way, ‘information’ helps to formulate hypotheses
that guide research and sharpen our understanding of what we
actually mean when we use the term. There is nothing wrong with
this strategy, particularly if it leads to testable, falsifiable predic-
tions. In fact, adopting such heuristic terms has a long and
continuing history in the biological sciences. ‘Gene’, ‘memory’,
‘mental map’, ‘auditory template’ and ‘neural representation’ are
other examples of words or phrases that scientists have used to
label an entity whose physical properties they are only beginning to
understand. The inability to specify precisely the information
conveyed by a vocalization (that is, its meaning to a listener) does
not prove that information is entirely absent.
The critics argue that we should abandon the concept of infor-
mation, but what programme of research do they offer in its place?
To reject the informational hypothesis entirely, we need evidence
that recipients in studies like those reviewed here acquire no
information. Although the ‘manipulative’ hypothesis has been
around for many years, no such data are available. The influence
that signallers have on receivers may help understand cases of
sensory exploitation (e.g. Ryan 1990), but years of research on this
topic do not compel us to abandon the concept of information.
Indeed, if there exist cases of sensory exploitation without
information transfer, the only way to make this distinction is to ask
if a signal contains information, a question the critics see as
‘ill-posed’. We agree that it is important to consider the form that
signals take, and can imagine some useful experiments to test
whether animals more readily learn pairings of certain types of
signals with particular outcomes, but this work would be fully
compatible with the informational perspective.
‘Teleological’ and ‘Circular’ Reasoning?
Both Owings & Morton (1998) and Rendall et al. (2009) object to
the ways in which scientists studying animal communication have
borrowed the term information from linguistics. Rendall et al.
(2009) believe that this approach is ‘both teleological and
circular’ because it ‘virtually guarantees’ that animal communica-
tion and language ‘will be “found” to be similar’ (page 238). In fact,
however, quite the opposite is true. The informational approach has
instead played a major role in uncovering differences between
species, and between animal communication and language, in the
kinds of information recipients acquire from signals.
For example, experiments on nonhuman primates have
repeatedly shown that, whereas human listeners acquire infor-
mation about a speaker’s mental states (intentions, beliefs, desires)
during conversation, nonhuman primate listeners appear to make
no such attributions (reviewed in Tomasello & Call 1997; Cheney &
Seyfarth 2007). Research on the information conveyed by vocali-
zations has thus led the way in revealing differences between
human language and nonhuman primate communication.
Curiously, Rendall et al. (2009) support their argument against
the ‘informational’ hypothesis by noting that recent studies have
revealed ‘an informational disconnection between signallers and
perceivers [suggesting] that they do not share the same represen-
tational parity that characterizes human speech’ (page 235). But
this conclusion, first proposed by Premack (1972, 1975) and later
elaborated (e.g. Cheney & Seyfarth 1990, 1996, 1998; Seyfarth &
Cheney 2003), emerged not as a result of their ‘manipulative’
approach but instead as a consequence of the very informational
perspective that they decry. Results demonstrate that, contrary to
the critics’ claims, the informational view is fully compatible with
distinct roles for signallers and receivers.
Finally, the lack of language-like intentionality in animal
communication does not mean that no continuities exist between
human and animal communication, nor should it prompt us to
abandon a research programme designed to search for precursors
of language in animal communication and cognition. Marler (1982)
was one of the first to examine similarities among birds, primates
and humans in meaning and the categorization of signals (see
also Cheney & Seyfarth 1990, 2007). Several recent studies have
documented psychological skills in nonhuman primates, such as
sensitivity to the presence of an audience or an awareness of the
directedness of a vocalization, that may well have been precursors
of intentional signalling during the course of human evolution (e.g.
Engh et al. 2006; Wich & deVries 2006). It would be impossible to
formulate hypotheses about the evolution of language if one started
with the premise that testing for language-like attributes in animal
communication is off-limits. Several recent studies in nonhuman
primate neurophysiology, inspired by work on primate vocaliza-
tions, demonstrate the value of a research programme that tests for
language-like processing in primates and other animals (e.g.
Ghazanfar & Santos 2004; Gifford et al. 2005; Belin 2006;
Ghazanfar et al. 2008; Petkov et al. 2008).
CONCLUSION
The critics have erected a straw man: most ethologists make no
claim that the signals they study are like language. The critics’ view
is also inconsistent with evolutionary theory. Recipients are not
powerless, unable to resist certain signals; instead, their responses
will evolve to reflect their own interests, and will depend on
both a signal’s physical properties and the information they acquire
from it.
Many studies argue against the critics’ hypothesis. They show
that calls with arousing or aversive features may also contain
 R.M. Seyfarth et al. / Animal Behaviour 80 (2010) 3e8
information that affects receivers’ responses; receivers’ responses
can be highly flexible; acoustically similar calls can elicit different
responses; acoustically different calls can elicit similar responses;
and ‘eavesdropping’ animals respond to the relationship instanti-
ated by signal sequences. All of these results support the view that
animal signals encode a surprisingly rich amount of information,
that the content of this information can be studied scientifically,
and that recipients’ responses depend at least in part on the
information they acquire from signals.
Far from being ‘teleological’ and ‘circular’, research inspired by
the informational perspective has clarified differences in the
mechanisms that underlie the behaviour of signallers and recipi-
ents; revealed differences between species in the information that
recipients acquire from signals; suggested fundamental differences
between language and animal communication; and inspired
a growing number of studies that examine the neurophysiological
basis of call meaning. The informational hypothesis thus continues
to prove its value in the most important way possible: by suggesting
observations and experiments that drive our field forward.
Acknowledgments
We thank Michael Beecher, Tabitha Price, William Searcy and
two anonymous referees for comments on the manuscript.
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